Genetic variation of Central European oaks: shaped by evolutionary factors and human intervention?

Oak species (Quercus spp.) in Central Europe grow on a relatively wide range of sites. Due to the economic importance of oak for its wood and other products, oak forests have long been managed by humans. This raises the question whether adaptation and/or human activities-especially the moving of propagules-have left their footprints on the genetic variation of oak populations. To address this question, we focused on the Upper Rhine Valley, a densely populated area today that was settled by humans early on. Here, the three most common native Central European oak species can be found. We studied their genetic variation across a large number of oak stands, growing on different sites and having different silvicultural histories, using neutral and EST-derived microsatellite markers. At the interspecific level, we showed that Quercus robur is relatively well delimited, while Quercus petraea and Quercus pubescens are more closely related. Natural hybridization might explain the increased genetic introgression between these two species. Within species, we found a low differentiation among populations of Q. robur and Q. petraea. In spite of forest fragmentation, we detected no spatial genetic barriers. However, we found that populations of Q. pubescens, a species with a marginal distribution in the study area were spatially structured. Genetic drift but also unidirectional introgressive hybridization with Q. petraea may account for this. Regarding the question of adaptation, we considered soil flooding, texture, drainage, and calcium carbonate in the upper horizons as physiologically important site condition variables. But with multivariate statistics, we could not find any significant effects of these parameters on genetic differentiation. Although there was no evidence for natural selection due to adaptation in stands of Q. robur, we demonstrated that age had a significant effect on their genetic variation and that stands established after the end of the Second World War had higher genetic diversity. We interpret these findings as being the result of an increase in large-scale transfers of reproductive materials during this time period and discuss arguments supporting this hypothesis. Finally, we consider the implications of these results for forest management.